Endocranial Morphology and Phylogeny of Palaeozoic Gnathostomes (Jawed Vertebrates)

Abstract: Gnathostomes, or jawed vertebrates, make up the overwhelming majority of modern vertebrate diversity. Among living vertebrates, they comprise the chondrichthyans (“cartilaginous fishes” such as sharks, skates, rays, chimaeras) and the osteichthyans (“bony fishes” or bony vertebrates, inclusive of tetrapods). Gnathostomes appear to have originated in the early Palaeozoic Era, but their early fossil record is fairly scant. The best fossils appear first in the Late Silurian and Devonian periods. Much of gnathostome diversity owes to unique adaptations in the internal skeleton of their head (the endocranium). The endocranium is composed of the braincase, jaws, hyoid arch, and branchial arches, which sometimes fossilise when they are composed of bone or calcified cartilage.The purpose of this thesis is to describe and compare the fossilised cranial endoskeletons of a variety of different Palaeozoic gnathostomes. The objective is to test current conceptions of gnathostome interrelationships (i.e. phylogeny) and infer aspects of key morphological transformations that took place during the evolution of Palaeozoic members of this group. Two key areas are examined: the morphology and interrelationships of Palaeozoic gnathostomes and the morphology of the visceral arches in sarcopterygian fishes.New data on the visceral arches are described from the stem tetrapods Panderichthys and rhizodontids. These provide insight into the sequence of character acquisition leading to the tetrapod middle ear. Panderichthys shows key features of the tetrapod middle ear chamber were established prior to the origin fo digited limbs. New morphological data are described from the “acanthodian” fish Ptomacanthus. Ptomacanthus provides only the second example of a well-preserved braincase from any member of this group. It shows dramatic differences from that of its counterpart, Acanthodes, providing new evidence for acanthodian paraphyly. New interpretations of basal gnathostome and osteichthyan phylogeny are presented, challenging or enriching existing views of these problems.